Bromalites from the Middle Triassic of Poland and the rise of the Mesozoic Marine Revolution

Palaeogeography, Palaeoclimatology, Palaeoecology 321–322 (2012) 142–150

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Bromalites from the Middle Triassic of Poland and the riseof the Mesozoic Marine Revolution

Mariusz A. Salamon a,⁎, Robert Niedźwiedzki b, Przemysław Gorzelak c, Rafał Lach a, Dawid Surmik a,c

a University of Silesia, Faculty of Earth Sciences, Department of Palaeontology and Biostratigraphy, Będzińska Str. 60, 41-200 Sosnowiec, Polandb Wrocław University, Institute of Geological Sciences, Cybulskiego 30, 50-204 Wrocław, Polandc Institute of Paleobiology, Polish Academy of Sciences, Twarda Str. 51/55, PL-00-818, Warsaw, Poland

⁎ Corresponding author.E-mail address: [email protected] (M.A. Sala

0031-0182/$ – see front matter © 2012 Elsevier B.V. Alldoi:10.1016/j.palaeo.2012.01.029

a b s t r a c t

Article history:Received 9 September 2011Received in revised form 17 January 2012Accepted 21 January 2012Available online 2 February 2012

Keywords:BromalitesRegurgitalitesMesozoic Marine RevolutionPredator–prey interactionsDurophagous predationEscalationTriassic

Durophagous predation has been an important cause of significant evolutionary changes in the history of life.One of the most dramatic predation-driven changes in marine ecosystems occurred during the middle andlateMesozoic, which has been called theMesozoicMarine Revolution (MMR). At this time, diversification of var-ious predators elicited many escalation-related adaptations among prey taxa including trends toward infaunali-zation, elaboration of armor shells, and environmental restriction. However, the rise of this phenomenon hasbeen the source of considerable debate. Initially, it has been argued that these major evolutionary changesbegan in the Jurassic and continued to accelerate in the Cretaceous. Although recent reports have shown thatthe MMR may have actually started soon after the end-Permian extinction in some groups, Triassic records ofpredation on benthic organisms are still very scant. Here, we report several bromalites (most probably regurgi-talites) from the Middle Triassic of the Gogolin Formation, Upper Silesia, southern Poland. They are in a form ofdistinct and packed accumulations of intermingled fossil remains composed primarily of various angular bivalveshell fragments with sharp, non-abraded margins as well as crinoid ossicles with common signs of breakage.These dispersions of material are up to 99 mm in maximum diameter. It is suggested that these accumulationsrepresent orally ejected waste produced by durophagous animals, most probably by durophagous sharks, colo-bodontid fish, placodonts, and some pachypleurosaurs or sauropterygian reptiles. All of these taxa have beenrecorded in the Middle Triassic of Poland. The feasibility of these vertebrates as potential agents of the presentbromalites is discussed. We suggest that many morphological and behavioral innovations in the Triassic gastro-pods, bivalves and crinoids are escalation-related adaptations to durophagous predators and that the MesozoicMarine Revolution was a far more prolonged evolutionary event than its name indicates.

© 2012 Elsevier B.V. All rights reserved.

1. Introduction

The Triassic was one of the turning points in the history of life. Fol-lowing the end-Permian extinction event that killed 80–90% of species(Benton and Twitchett, 2003; Knoll et al., 2007), recovery of benthicmarine communities took place during the Early Triassic (e.g.Twitchett et al., 2004; Hofmann et al., 2011) or early Middle Triassic(e.g. Hu et al., 2011) leading to the rapid expansion through previouslyrealized morphospace as well as many morphological and behavioralinnovations (Foote, 1995; Baumiller et al., 2010). However, debatecontinues regarding the importance of biotic versus abiotic factors inthe evolution of Triassic communities. One hypothesis implies thatthe varied morphological and behavioral features of Triassic benthicinvertebrates, such as gastropods, bivalves and crinoids, may havebeen driven by escalating predator–prey interactions (e.g. Stanley,1968; Nützel, 2002; Hautmann, 2004; Baumiller et al., 2010). In

mon).

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contrast, McRoberts (2001), for example, suggested that many ofthese so-called predatory adaptations in bivalves were in fact pread-aptations, unrelated to coevolving predatory groups, due to severalinterconnected abiotic and biotic causes associated with the recoveryafter the end Permian mass extinction. The latter author argued thatdespite the fact that a number of durophagous predatory taxa werepresent during the Triassic, they were not widespread and/or abun-dant enough to cause a marked selection pressure on their victims.

Although various direct and indirect indicators of predation areavailable to paleontologists, the fossil record of Triassic predation israther scanty, and its incompleteness leaves many open questions. Asyet, only a few reports on boring and crushing predation on mollusksare known from the earlyMesozoic (Tintori, 1995; Harper, 2006 and lit-erature cited herein). Evidence of predation on Triassic crinoids is alsoscarce and comes from the specimens yielding regeneration traces(e.g., Weissmüller, 1998; Oji, 2001) or signs of bite marks (Baumilleret al., 2010).

In recent marine environments fragmentation of the skeletons ofbenthic invertebrates, such as bivalves, caused by the shell-crushingpredators is very common (Cate and Evans, 1994 and literature cited

Fig. 1.Map of Poland with investigated area indicated and enlargement of Upper Silesiawith the sampled Wojkowice quarry indicated (circle).Taken from Zatoń et al. (2008) and authors cited therein; slightly modified.

143M.A. Salamon et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 321–322 (2012) 142–150

herein). It has been generally argued that extant birds, sharks and somefish may regurgitate (or vomit) food and difficult-to-digest food resi-dues that accumulate in their stomach (e.g. Strassen, 1914; MacGinitieand MacGinitie, 1949; Müller, 1957; Cadée, 1994). However, differenti-ation between shell-fragmentation caused by regurgitating duropha-gous predators versus physical factors is difficult in the fossil record.Recently, Oji et al. (2003) suggested that incomplete mollusks with an-gular margins can be considered as a good proxy for durophagous pre-dation. Their assumption was based on neontological tumblingexperiments as well as quantitative analysis of shell fragments from anumber of Mesozoic and Cenozoic beds in Japan. Their results showedthat majority of shells from their tumbling experiments were abradedand only infrequently broken into angular fragments. By contrast, theynoted that accumulations with angular and non-abraded shell frag-ments increased during the early Cenozoic, in the contemporaneous di-versification of durophagous teleost fish and decapod crustaceans. Thelatter authors therefore concluded that the majority of these accumula-tions in the studied bedswere the result of durophagous predation. Sub-sequently, Zatoń and Salamon (2008) found very similar fossilaccumulations with angular and non-abraded shell fragments from theMiddle Jurassic of Poland that were ascribed to durophagous predation.

The main goal of the present paper is to provide evidence of thistype of predation in the Middle Triassic and to examine the evolution-ary history of Triassic gastropods, bivalves and crinoids in the contextof evolving durophagous predators.

2. Geological setting, materials and methods

Field work has been carried out in “Wojkowice” abandoned quarrylocated in the southern part of Silesian-Cracow Monocline in UpperSilesia, southern Poland (Fig. 1). The Triassic sediments of this quarrywere deposited in the Germanic Basin on the northern margin of the

Fig. 2. Stratigraphical section of the northern part (N) and southern part (S) of theWojko-wice Quarry. 1— dolomitic limestones; 2— cellular dolomitic limestones; 3— organodetri-tal limestones with bivalve detritus and columnals; 4 — marly limestones; 5 — pelliticlimestones with abundant shells of bivalves; 6— pellitic limestones; 7—wavy limestones;8 — nodular limestones; 9 — vertebrate remains; 10— Dadocrinus columnals; 11— encri-nids columnals; 12 — intraclasts; 13 — bromalites; 14 — Rhizocorallium commune; 15 —

numerous gastropods; 16 — numerous Plagiostoma; 17 — numerous Pseudocorbula sp.;18 — numerous Gervillia sp.; 19 — Thalassinoides; 20 — Holocrinus columnals; I — Roetian;II—“limestoneswith Entolium andDadocrinusunit”; III—“firstwavy limestones unit”; IV—

“cellular limestones unit”; V — “thick-bedded limestones” and “wavy limestones unit”.

Tethys Ocean (e.g., Szulc, 2000). In Wojkowice, dolomitic limestonesand dolomites of the Upper Buntsandstein (Röt) as well as limestonesof the lowermost part of the Lower Muschelkalk (Gogolin Formation

Fig. 3. Pie charts showing faunal contents (A1, B1, C1, D1) and size-frequency distribu-tion of the bivalve shell fragments (A2, B2, C2, D2) in three dissolved regurgitalite accu-mulations (A, B, C) and in surrounding rock sample outside accumulations(D) collected in the Wojkowice Quarry, southern Poland, Gogolin Fm., “limestoneswith Entolium and Dadocrinus unit”, Upper Olenekian/Lower Anisian. A sample no:W1. B sample no.: W5. C sample no.: W6.

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sensu Kowal-Linka, 2008) are exposed (Fig. 2). In the lower partof the Gogolin Formation, the “limestones of the Entolium andDadocrinus unit” (sensu Assmann, 1944) crop out, represented byorganodetrital and crinoidal limestones with numerous Dadocrinuscrinoid ossicles and bivalve shells of Entolium discites. Withinthis unit, 16 distinct fossil accumulations were found. They occurwithin fossiliferous limestone layer containing numerous undamagedE. discites shells, Dadocrinus ossicles, undiagnostic gastropods andbones of marine reptiles, fish scales (e.g., Gyrolepis) and shark teeth(Acrodus, Palaeobates). Shells of a bivalve Plagiostoma are extremelyrare here. Contrary to some beds of fossiliferous limestones above,intraclasts occur rarely in this part of the section. The layer with dis-tinct fossil accumulations is capped by thin (a few mm thick) marl-stones with numerous external molds of small gastropods.

Based on magnetostratigraphic studies of the Röt andMuschelkalkin Upper Silesia (Nawrocki and Szulc, 2000), the Röt and the lower-most part of the “limestones with Entolium and Dadocrinus unit”should be included into the late Olenekian, whereas the upper partof the “limestones with Entolium and Dadocrinus unit” representsthe Aegean. It has been argued that the sediments of the Röt were de-posited in a sabkha environment with marine incursions whereas the“limestones with Entolium and Dadocrinus unit” were formed in shal-low water and open marine conditions (Bodzioch, 1991). Beds of fos-siliferous limestones with Entolium and Dadocrinus in Upper Silesiaare generally interpreted as proximal tempestites (Hagdorn andSzulc, 2007a, b). The layer with distinct fossil accumulations differsfrommost similar beds of fossiliferous limestones described in the lit-erature (e.g., in Hagdorn and Szulc, 2007a, b) because of a bimodalityof the state of preservation of fossils, i.e., fossil remains in distinct ac-cumulations are commonly preserved as fragments of shells and iso-lated crinoid ossicles with signs of breakage, whereas the same fossilsoutside these distinct accumulations are preserved as commonlyundamaged bivalve shells or relatively well-preserved crinoid pluri/columnals.

All distinct accumulations were photographed, and the main fea-tures such as shape, size and faunal content were noted (Fig. 3,Table 1). The next step was to transport large slabs with accumula-tions to the laboratory. Initially, all samples were carefully cleanedin hot water. Later the samples were examined in more detail.Three accumulations were selected for more detailed taphonomic ob-servations (for methodology see Zatoń and Salamon, 2008; accumula-tions no. W1, W5 andW6; see Fig. 3). Selected slabs were cut to leaveonly characteristic fossil accumulations and were heated in waterwith perhydrol (a 30% aqueous solution of hydrogen peroxide) upto 100 °C for a maximum 20 min. The boiling procedure was repeatedup to as many as 6 to 9 times. The resulting residue was carefullycleaned and dried. All bivalve shell fragments that were the basic fau-nal component, were measured by dry sieving using a series of sieveswith mesh sizes ranging from 1.25 to 20 mm. All fragments from aparticular mesh range (20–10, 10–5, 5–3.15, 3.15–2.5, 2.5–1.25 mm)were counted. Basic taphonomic features [such as signs of abrasion,dissolution, bioerosion, and signs of breakage (in case of crinoids)]were also investigated (Table 2).

Spot elemental analyses on a polished and carbon-coated frag-ments of the rock matrix that infilled the selected fossil accumula-tions (GIUS 7-3591G7;8) were performed with a scanningmicroscope Philips XL — 20 coupled with an EDS detector ECON 6(system EDX — DX4i) at the Institute of Paleobiology, Warsaw,Poland. The material is housed in the Faculty of Earth Sciences, Uni-versity of Silesia, Sosnowiec, under catalogue number: GIUS 7-3591.

3. Results

All investigated accumulations were clearly distinct from the sur-rounding sediment and have an elliptical or subcircular outline(Figs. 4–7). Their dimension varies from about 13 mm–18 mm to

about 55 mm–99 mm (Table 1). All accumulations were separated,but some of them were densely grouped (sample W3 and W7; seeTable 2). In all of the accumulations, the fossils were represented byonly four fossil groups (Fig. 3, Table 1). All fossils were intermingledand distributed from the center to the margin.

As mentioned above, in all accumulations only four taxonomicgroups occurred. Most fossils were bivalve shell fragments (almost83% of all fossil remains) that possessed sharp and angular edges.Among bivalves, thin shell fragments with indistinct fine ribs(probably Entolium) predominate (up to about 69% of all bivalvedetritus), whereas thick shell fragments with coarse ribs (possiblyPlagiostoma) are rarer. Apart from bivalve shells, crinoid columnalsand brachials (Dadocrinus sp.) with common signs of breakage(as much 45% in a sample, see Table 2) were present, as well asfragmented gastropods of unknown affinities and three teeth ofmarine reptiles (Fig. 3, Table 1).

Taking into account the total amount of bioclasts (mostly bivalvefragments) from a particular fraction from selected samples, it is evi-dent that the 5.0 mm–3.15 mm size fraction and larger predominates

Table 1The general features of accumulations collected in the Wojkowice Quarry, southern Po-land, Gogolin Fm., “limestones with Entolium and Dadocrinus unit”, Upper Olenekian/Lower Anisian.

Sample(regurgitalites) number

Max. size(given in mm)

Shape Fossils occurred in theregurgitalites

W1 53×39 Ellipsoidal Bivalves, crinoids,gastropods, vertebrates

W2 51×37 Oval BivalvesW3/A 52×42 Ellipsoidal BivalvesW3/B 63×34 Ellipsoidal Bivalves, crinoidsW3/C 48×46 Oval BivalvesW3/D 18×13 Oval BivalvesW4 42×38 Oval Bivalves, crinoidsW5 52×20 Ellipsoidal Bivalves, crinoidsW6 37×32 Oval Bivalves, crinoidsW7/A 65×54 Ellipsoidal BivalvesW7/B 34×24 Ellipsoidal Bivalves, crinoidsW7/C 42×31 Ellipsoidal Bivalves, crinoidsW7/D 39×31 Ellipsoidal BivalvesW7/E 38×22 Ellipsoidal Bivalves, crinoidsW7/F 25×16 Ellipsoidal Bivalves, gastropodsW7/G 99×55 Ellipsoidal Bivalves, crinoids

Fig. 4. A, Investigated level of the “limestones with Entolium and Dadocrinus unit”(Lower Gogolin Beds, Upper Silesia, southern Poland, southern wall of the quarry)with in situ bromalites (regurgitalites?) accumulations (dotted lines) of fossils remainsfrom (B–D (enlargements)). D2, line drawing of fossil remains from accumulation D. Acoin as scale bar for (B, D) (diameter equals 1.85 cm), hammer for A scale bar for Cequals 10mm. Acronyms: B GIUS 7-3591, C GIUS 7-3591B, D GIUS 7-3591D.

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(Fig. 1). The sizes below 3.15 mm in three investigated accumulationsare rare. However, samples no. W1 and W6 had a significant amountof size fraction 2.5 mm–1.25 mm (Fig. 3).

A very small percentage of the fossils in the accumulations havesigns of bioerosion, abrasion or dissolution (Table 2). The abrasion isdocumented only on a few shell fragments of samples W1 and W5.The encrustation by epibionts and dissolution were not recorded infossils from accumulations.

Our spot geochemical analysis of fragments of the rock matrix(not shell) that infilled the two selected fossil accumulations (GIUS7-3591G7 and GIUS 7-3591G8) revealed them as calcium carbonate[C: 12.62 wt%; O: wt% 50.68; Ca: wt% 36.7 (first sample) and C:12.9 wt%; O: wt% 49.17; Al: 0.16 wt%; Si: 0.14 wt%; Mg: 0.37 wt%;Ca: 36.7 wt% (second sample); average values obtained from 3 spots].

4. Discussion

4.1. What was the culprit?

Considering the origin of fossil accumulations described here, sev-eral explanations should be taken into account: 1) episodic accumula-tion of fossil debris due to waves or currents, 2) detached load casts(pseudonodules) which sank into the bottom, 3) intraclasts, 4) com-paction influence, 5) infills of decapod burrows, 6) coprolites or 7)regurgitalites (regurgitates or vomites).

Sedimentation by currents or waves is excluded, as such depositsare typically represented by rounded and abraded shells that form ac-cumulations which are commonly parallel to the shoreline. By con-trast, the present accumulations are randomly oriented, and theyare composed of angular, sharp and unabraded shell fragments ofvarying size (from 1.25 to 20 mm, e.g. sample W1 in Fig. 3), whichsuggests that sorting of these bioclasts was not involved.

Explanation of these accumulations as load casts is also unlikelybecause the investigated layer is overlain by an undisturbed thin

Table 2The general features of dissolved accumulations collected in the Wojkowice Quarry,southern Poland, Gogolin Fm., “limestones with Entolium and Dadocrinus unit”, UpperOlenekian/Lower Anisian.

Sample no: W1 Sample no: W5 Sample no: W6

Total number of remains 186 47 62Abrasion (bivalves) 6% 4% 2%Bioerosion (bivalves) 1% 2% 1%Breakage (crinoids) 33.3% 16.7% 45%

Fig. 5. A, Investigated level of the “limestones with Entolium and Dadocrinus unit”(Lower Gogolin Beds, Upper Silesia, southern Poland, southwestern wall of the quarry)with in situ bromalites (regurgitalites?) accumulations (dotted lines) of fossils remains(arrows) from (B–C). A coin as scale bar for C (diameter equals 1.85 cm), hammer forA; scale bar for B equals 10 mm. Acronyms: B GIUS 7-3591E, C GIUS 7-3591F.

Fig. 6. A, Investigated level of the “limestones with Entolium and Dadocrinus unit”(Lower Gogolin Beds, Upper Silesia, southern Poland, southeastern wall of the quarry)with in situ bromalites (regurgitalites?) accumulations (dotted lines) of fossils remainsfrom (B–E). A coin as scale bar for C–E (diameter equals 1.85 cm), hammer for (A, B).Acronym: B GIUS 7-3591G.

Fig. 7. A and B, Enlargements of selected bromalites (regurgitalites?) from the “lime-stones with Entolium and Dadocrinus unit” (Lower Gogolin Beds, Upper Silesia, south-ern Poland). Scale bar 10 mm B. C, Crinoid ossicles bearing signs of breakageobtained from bromalites (regurgitalites?) no. W1. Scale bar 1 mm.

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marlstone bed that is capped by a similar layer of limestones withidentical faunal taxa as the bed with these distinct fossil accumula-tions. Additionally, the lower surface of this layer is straight withoutany evidence of load casts. The rock surrounding the distinct accumu-lations lacks intraclasts but contains numerous undamaged shells orlarge fragments of thin-shelled bivalves Entoliumwhich are indicativeof a low energy environment. Furthermore, our accumulations do nothave sharp margins which are typical for intraclasts.

In the Lower Muschelkalk decapod burrows are common. Thalassi-noides is common in Mesozoic sediments and the walls of some tun-nels are lined with biodebris (such as shell fragments or fish scales)(Bromley and Frey, 1974). Additionally, numerous burrows of Thalas-sinoides are passively filled (e.g. Kamola, 1984). Therefore, the tunnelssometimes contain coarser material, whereas the background is com-posed of pelitic sediment (e.g. Kędzierski and Uchman, 2001). How-ever, the accumulations on the bedding surface of the GogolinFormation are more or less elliptical and do not penetrate deep intothe sediment, in contrast to the decapod burrows. Furthermore, inthe Lower Gogolin Beds of Wojkowice, the ichnofossil Thalassinoideswas not recorded. Other observed ichnofossils which commonlyoccur in the beds under study are Rhizocorallium, which do not con-tain any fossil debris and are only present in the pelitic sediments.

It is possible that compaction could have been involved in the pro-duction of the presently observed accumulations (Vermeij, 2002;Zuschin et al., 2003). However, the fragmentation pattern of shellsisolated differs significantly from those resulting from compaction.They often occur as isolated single valves of different individuals,

whereas fragmented parts of a shell produced by compaction arestill close together. Furthermore, the influence of compaction is un-likely as only specimens in distinct fossil accumulations are crushed.

Both taphonomic features (evidence of angular shell fragmentswithlarge size dispersionwith sharp, non-abradedmargins and crinoid ossi-cles with common signs of breakage) and the overall sedimentologicalcontext imply biomechanical production. The most probable explana-tion is that they represent coprolites or regurgitated remains producedby durophagous predators. The fact that shell fragments of possible Pla-giostoma are much more common in the accumulations than in sur-rounding sediment is consistent with this interpretation and maysuggest selection by a predator. Furthermore, fossils in accumulationsand in surrounding sediment are differently preserved, which stronglysupports biological origin. Indeed, these accumulations strongly resem-ble those retrieved from the digestive tracts of the durophagous fishPogonias by Cate and Evans (1994) as well as from the Late Triassic ofItaly (Tintori, 1995) and Jurassic of Poland (Zatoń and Salamon, 2008).

The most common definition of ‘coprolite’ is fossilized animal dung.On the other hand, regurgitalites (=regurgitates or vomites sensuWood, 1980) represent the expulsive material from the mouth, pharynx,or esophagus, usually characterized by the presence of undigested food.However, the terminology relating to coprolites and other fossilized prod-ucts of digestion is rather poorly defined. There are several features, how-ever, that allow recognition of coprolites and regurgitalites. According toPollard (1990) and Chin (2002), the coprolites are mostly composed ofcalcium phosphate, therefore differ from the accumulation in discussionthat is composed of calcium carbonate (see Results section). During dia-genesis, however, the coprolite mass is sometimes replacedwith siderite,limonite or silica. Pollard (1990) argued that coprolites retain featuressuch as spiral foldings or vascular markings. Hattin (1996) distinguishedfive different coprolites that are formed by carbonate fluorapatite andhave cylindrical shapes. In comparison to coprolites, regurgitalites gener-ally lack a phosphatic matrix and contain better-preserved and recogniz-able food remains that have very often non-abraded margins (see alsoZatoń and Salamon, 2008). Furthermore, in regurgitalites, skeletal re-mains are less affected by the digestive processes.

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Hunt (1992) proposed the term ‘bromalite’ to encompass all fos-silized products of digestion. Other terms used by Hunt (1992) andNorthwood (2005) were: 1) cololites: non-valvular intestinal con-tents; 2) coprolites: solid dietary waste expelled through the anusor cloaca; 3) enterospirae: contents preserved in the valvular intes-tine; 4) gastrolites: fossilized stomach contents; 5) regurgitalites:orally ejected waste (this term is adopted here).

4.2. Durophagous predators in the Middle Triassic of the Germanic Basin

Based on dimensions of fossil accumulations (up to about 99 mm),it is evident that most of themwere not produced by predatory inver-tebrate(s) (e.g. echinoids, asteroids or decapods) but rather by largerdurophagous vertebrate(s). It is difficult to assign the present broma-lites to specific producer other than a durophagous predator.

In the lower part of the Gogolin Formation from Upper Silesia nu-merous remains of durophagous predators were recorded. Amongthem, broad, rather low and blunt crushing teeth without sharp cus-plets are the most common, i.e. shark teeth of Acrodus, Lissodus andPalaeobates (e.g., Liszkowski, 1993) and colobodontids (actinoptery-gians) teeth. Indeed, fossil regurgitalites of Late Cretaceous duropha-gous sharks have been documented and they are very similar in thesize and shape to those documented in the present study ((Hattin,1996, Fig. 8; compare also shark regurgitalites from the Pennsylva-nian of Indiana (USA) (pl. 44 C–D, pl. 49 A; Zangerl and Richardson,1963)). Other potentially molluscivorous vertebrates (Fig. 8A–D),which are relatively common in the investigated formation of UpperSilesia, are reptiles, especially placodonts (Surmik, 2010; see alsoFig. 8D), pachypleurosaurs (Chudzikiewicz, 1983) and sauropterygianreptiles with uncertain taxonomical affinity (e.g., Hemilopas mentzeli).Although some placodonts are generally considered as typical duro-phagous predators (Rieppel, 2002; Walker and Brett, 2002; Motani,2009), recently Diedrich (2010, 2011), based on functional morphol-ogy of their jaws and teeth, postulated that they were feeding on ma-rine algae. However, Yiang et al. (2008) described a Triassic placodontfrom China with exceptionally preserved stomach content consistingof bivalves. Pachypleurosaur remains from the Upper Silesia are com-monly preserved as isolated and incomplete fragmentary bones, thusthis group in the Silesian Muschelkalk is rather poorly known. How-ever, in the Röt/Lowermost Muschelkalk (Lower Anisian) of

Fig. 8. Vertebrate durophagous remains occurring in Upper Silesia of Poland. A, Palatineteeth of Colobodus (Actinopterygii) from Lower Muschelkalk (Lower Anisian), NakłoŚląskie, near Tarnowskie Góry, WNoZ/S/7/141. B, Jaw of enigmatic fish Nephrotus chor-zowiensis, Lower Muschelkalk, Zakrzów near Gogolin, housed in Museum of DepositsGeology, Gliwice Silesian University of Technology. C, Jaw fragment of durophagouspredator probably ichthyopterygian from Röt/Muschelkalk boundary, Rogoźnik Quar-ry, near Wojkowice, WNoZ/S/7/68. D, Placondontid dentale fragment from LowerMuschelkalk, Krapkowice Quarry, near Gogolin, housed in Museum of Deposits Geolo-gy, Gliwice Silesian University of Technology. Scale bars 10 mm.

Winterswijk (Netherlands) and in the upper part of the LowerMuschelkalk near Freyburg remains of pachypleurosaur belongingto Anarosaurus heterodontus are very common (Hagdorn andRieppel, 1998; Őosterink et al., 2003; Klein, 2010). Its jaw structurestrongly indicates that it was able to prey on taxa protected by hardskeleton, such as bivalves (Klein, 2009). Teeth of H. mentzeli were de-scribed from the lower part of the Gogolin Formation (Rieppel, 1995),in the Lower Muschelkalk from Chorzów (Schmidt, 1928). They arealso known from the old collection of the Department of Palaeozool-ogy of University of Wrocław collected at Zakrzów near Gogolin. Inthe lower part of the Lower Muschelkalk of the Germanic Basin(Maisch and Matzke, 2001) remains of ichthyosaurs Phalarodonwith crushing teeth occur rarely. According to Massare andCallaway (1990), this ichthyosaur was feeding on mollusks.

4.3. Early Mesozoic Marine Revolution

The term “Mesozoic Marine Revolution” (MMR) introduced byVermeij (1977), is given to the marked changes (such as increasedinfaunalization, shell sturdiness, environmental restriction) of shal-low benthic communities due to radiation of durophagous and dril-ling predators. It was suggested that MMR started in the Jurassicand continued at an accelerated pace during the Cretaceous(Vermeij, 1977). McRoberts (2001) stressed that durophagous preda-tors had low abundances and limited distribution during the Triassic.Additionally, Kowalewski et al. (1998) pointed out that predatorydrilling was relatively ineffective and largely lost during the Triassic.

However, it has been shown that many taxa, such as: ceratites,Chondrichthyes (hybodontids), some ichthyopterygian and saurop-terygian clade (including: placodonts, pachypleurosaurs, some pisto-saurs and nothosaurs) also developed crushing abilities in the Triassic(Walker and Brett, 2002; Underwood, 2006; Kriwet et al., 2009). Newexamination of the Decapoda suggests that innovations for duro-phagy were also more broadly distributed in this group during theTriassic (Schweitzer and Feldmann, 2010). This suggests that Triassicpredators might have caused a significant selection pressure on theprey taxa. Indeed, Walker and Brett (2002) stressed that MMR com-prises a series of revolutions, starting during the Triassic. Recently,Vermeij (2008, 2011) concluded that the major evolutionary innova-tions among benthic fauna actually occurred during two periods, theLate Triassic to Early Jurassic and the Late Cretaceous.

In the case of mollusks, the marked escalation probably began nolater than the Carnian stage (Late Triassic) (Harper, 2006). Duringthis time, many macroevolutionary trends that can be interpreted asadaptive consequences of durophagous predation have been recog-nized. For example, Nützel (2002) observed that reticulate sculpturein gastropods became very common during the Carnian. The latter au-thor interpreted reticulate sculpture with beads, knobs, or spines thatmay increase shell strength, as a defensive adaptation. Recent studyby Kosnik et al. (2011) showed that an increase in mean gastropodreinforcement might appear much earlier, in the early Triassic.Hautmann (2004) pointed out that several independent clades ofcementing bivalves with new types of ligaments (alivincular-arcuateand alivincular-fossate) appeared during the Late Triassic. Experi-mental study by Harper (1991) has shown that cementation strate-gies in bivalves affords protection against predators such asasteroids and crabs. Furthermore, some Late Triassic bivalves devel-oped an alivincular-alate ligament type that allowed opening thevalves, and consequently effective “swimming”. Stanley (1977)drew attention to the fact that many infaunal clades of bivalves be-came common during the Triassic. This infaunal mode of life mayhave been particularly an effective protection from grazing predators(Walker and Brett, 2002). Also notable is a general increase in bivalveshell size during the late Triassic (Kosnik et al., 2011).

Not so long ago, it was assumed that predation on echinodermswas not intense during the Triassic (Schneider, 1988). This claim

148 M.A. Salamon et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 321–322 (2012) 142–150

was supported by Zatoń et al. (2008) who recorded extremely lowfrequency of arm regeneration in Middle Triassic ophiuroids. Howev-er, in a novel study, Baumiller et al. (2010) based on Recent in-situobservations and simulated experiments, showed that cidaroid seaurchins feed on live stalked crinoids, leaving distinct bite marks ontheir skeletal elements. Simultaneously, their taphonomic survey ofthe Early–Middle Triassic crinoid ossicles from Poland revealed thatmany of them bear similar bite marks. Thus, the latter authors inter-preted that the Triassic crinoids must have been subject to significantechinoid predation pressure that led to the major evolutionary radia-tion among crinoids during the Middle–Late Triassic. They listed sev-eral taxa with distinct morphological and behavioral novelties with amobile life habit allowing protection from benthic predators, such as:planktonic microcrinoids (roveacrinids), pseudoplanktonic stalkedcrinoids (traumatocrinids), benthic isocrinids capable of rapid crawl-ing, the primitive free-moving crinoids (paracomatulids) and pseudo-planktonic pentacrinitids. Although predation by echinoids and itsconsequences have received the most attention, sparse data on re-generation traces (Weissmüller, 1998; Hagdorn, 2011) as well asour findings of bromalites, imply that the Middle Triassic crinoidsmight have also been the prey of nektonic predators. The common oc-currence of the presently described bromalites (16 bromalites perabout 10 m2 of the bedding surface of the fossil bed, number compa-rable to those reported in some Cenozoic beds of Japan (Oji et al.,2003)) suggests that durophagous predation by nektonic predatorshas been intense during the Middle Triassic. It is important to notethat Oji et al. (2003) in their analysis of shell fragments in a numberof Mesozoic and Cenozoic beds in Japan demonstrated that angularand non-abraded shell fragments did not occur in the two early Trias-sic fossiliferous formations. Therefore, by a global extrapolation wemay speculate that a considerable increase in regurgitate accumula-tions in shell beds across the Early/Middle Triassic occurred. Althoughsuch extrapolations are risky, these data are coincident with the in-creased diversity of shell-crushing predators in the Middle Triassic(Harper, 2006, Fig. 3). Further study of bromalites from the early Me-sozoic deposits of Europe is planned to test this hypothesis.

To explore the possible selection pressure of nektonic predatorson the Triassic crinoids, we surveyed the literature for various

Table 3Triassic innovations in gastropods, bivalves and crinoids and their times of first appearanceData are taken from: Nützel (2002); Hautmann (2004); Wolkenstein et al. (2006); VermeijMessing (2011); Niedźwiedzki et al. (2011).

Innovation Time

GastropodsReticulate sculpture in gastropods Late T

BivalvesAlivincular-arcuate ligament type in cementing bivalves (Ostreidae) Late TAlivincular-fossate ligament type in cementing bivalves(Dimyidae, Plicatulidae, Spondylidae)

Late T(Carn

Alivincular-alate ligament type in “swimming” bivalves(Entoliidae, Pectinidae)

Late TRhaet

Obligately deep-boring bivalves Late TCrinoids

Autotomy (shedding) planes in the stalk and arms of Holocrinida and Isocrinida MiddlMuscular articulations allowing rapid crawling in Holocrinida and Isocrinida MiddlSpines or tubercles on latera in Encrinida and Millericrinida MiddlPurple to violet pigmentation in Encrinida MiddlAbility to regenerate the holdfasts in Millericrinida MiddlAxillaries prolonged into long aboral spines and laterally enlarged cirralsfor protection of the arms in Ainigmatocrinidae

Middl

Pseudoplanktonic lifestyles in Traumatocrinidae MiddlEleutherozoic lifestyle by stem reduction in Roveacrinida Middl

Miniaturization in Roveacrinida MiddlDeeper settings residence and strongly specialized morphologyin Cyrtocrinida

Late T

Pseudoplanktonic lifestyles in Pentacrinidae Late TEleutherozoic lifestyle by stem reduction in protocomatulids(Paracomatula and Eocomatula)

Late T

innovations that could be interpreted as antipredatory adaptations.In addition to five major innovations interpreted by Baumiller et al.(2010), we identified eight additional innovations that potentiallystimulated or reflected adaptation against nektonic predation(Table 3).

Taxa with possible anti-predatory innovations included: 1) Ani-sian holocrinids (Holocrinida) and isocrinids (Isocrinida) with autot-omy (shedding) planes (so-called cryptosyzygies in the brachials andsmooth cryptosymplectial facets in nodals) that are known to reducedamage and stalk/arm loss (e.g., Oji and Okamoto, 1994), 2) Anisianencrinids (Encrinida) and millericrinids (Millericrinida) with spines(Stiller, 2000; Dynowski and Nebelsick, 2011), 3) Anisian encrinidswith purple to violet pigmentation (e.g. Wolkenstein et al., 2006),4) Ladinian ainigmatocrinids (Ainigmatocrinidae) with axillaries pro-longed into long aboral spines and laterally enlarged cirrals, both forprotection of the arms (Hagdorn, 1988; Hagdorn and Rieppel,1998), 5–6) unstalked crinoids with eleutherozoic lifestyle (such asLadinian roveacrinids (Roveacrinida) and Norian-Rhaetian comatu-lids (Comatulida) (Hagdorn and Campbell, 1993; Hess and Messing,2011), 7) Ladinian planktonic “microcrinoids” (roveacrinids, Rovea-crinida) with miniature size (Hess and Messing, 2011), (8) Car-nian?/Rhaetian small stalked crinoids, cyrtocrinids (Cyrtocrinida)with strongly specialized morphology (specific structure of crownand arms that could be enrolled in a cavity formed by large medianprolongation of the second primibrachials or by interradial processesof the radials) and behavioral features (deeper habitats) (see Salamonet al., 2009).

5. Conclusions

Although recent reports indicated that innovations for durophagywere more broadly distributed during the early Mesozoic and amonggroups that might have even caused a significant selection pressureon the benthic communities, the Triassic record of durophagous pre-dation has been rather scanty. Discovery of frequent bromalites in theMiddle Triassic of Poland constitutes important evidence of predator–prey interaction in the early Mesozoic. We suggest that many mor-phological and behavioral innovations in the Triassic gastropods,

.(2008 and references therein), Baumiller et al. (2010 and references therein), Hess and

of origin Innovation category

riassic (Carnian) Defensive against nektonic and benthic? predation

riassic (Carnian) Defensive against benthic and nektonic predationriassician and early Jurassic)

Defensive against benthic and nektonic predation

riassic (Norian-ian)

Locomotion, defensive against benthicand nektonic predation

riassic (Carnian?) Habitat, defensive against benthic and nektonic predation

e Triassic (Anisian) Defensive against benthic and nektonic predatione Triassic (Anisian) Locomotion, defensive against benthic predatione Triassic (Anisian) Defensive against nektonic and benthic? predatione Triassic (Anisian) Defensive against nektonic predatione Triassic (Anisian) Defensive against benthic predatione Triassic (Ladinian) Defensive against nektonic predation

e Triassic (Ladinian) Habitat, defensive against benthic predatione Triassic (Ladinian) Habitat, locomotion, defensive against benthic

and nektonic predatione Triassic (Ladinian) Defensive against nektonic predationriassic (Carnian) Habitat, defensive against nektonic predation

riassic (Norian) Habitat, defensive against benthic predationriassic (Norian-Rhaetian) Habitat, locomotion, defensive against benthic

and nektonic predation

149M.A. Salamon et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 321–322 (2012) 142–150

bivalves and crinoids can be regarded as escalation-related-adaptations to durophagous predation (both benthic and nektonic)rather than adaptations unrelated to coevolving predatory groups assuggested by McRoberts (2001). Consequently, our data imply thatthe predation-driven Mesozoic Marine Revolution had alreadystarted during the Anisian and was a far more gradual evolutionaryevent than previously thought.

Acknowledgments

We would like to specially thank T. K. Baumiller (University ofMichigan) and W. I. Ausich (The Ohio State University) for many use-ful remarks and suggestions for syntactical improvement of the ms.Comments of three anonymous reviewers and the Editor, Dave Bott-jer, helped improved the manuscript. The junior author (RL) wouldlike to thank the UPGOW project for financial support. This projectwas supported by the National Science Centre grant no. UMO-2011/01/B/ST10/02639.

Appendix A. Supplementary data

Supplementary data associated with this article can be found inthe online version, at doi:10.1016/j.palaeo.2012.01.029. These datainclude Google map of the most important areas described in thisarticle.

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